The distribution,abundance, and the effects of fire on mound building termites (Trinervitermes and Cubitermes spp., Isoptera: Termitidae) in northern guinea savanna West Africa

Summary Termite mound densities in typical guinea savanna, Detarium, and grassland (boval) habitats in northern guinea savanna were determined by random quadratting of 2–3 sites in each habitat (100, 10x10 m quadrats per habitat). Dominant species in guinea savanna were T. geminatus (46 mounds ha^-1) and T. oeconomus (21 mounds ha^-1), in Detarium T. geminatus (59 mounds ha^-1) and C. curtatus (45 mounds ha^-1) and in boval C. curtatus (72 mounds ha^-1) and T. geminatus (22 mounds ha^-1). Only C. curtatus densities and total densities differed significantly between sites within habitats, but all species differed significantly in abundance between habitats. The composition of each community was related to general environment but no particular environmental variable was shown to be a major determinant of termite distribution. Evidence for the limitation of termite populations was obtained from indirect evidence of competition between colonies in Detarium, and by experimental manipulation of fire regimes in the typical guinea savanna habitat. Harvester termites increased four-five fold over two years in fire-protected plots as a result of increased food supplies. Total termite densities in the fire-protected community equilibrated to the new population density (100 mounds ha^-1) after only two-three years.     

Community response to experimental soil disturbance in a midsuccessional,abandoned pasture

To determine whether soil disturbance by digging and burrowing mammals altered community structure and the rate of succession in a midsuccessional abandoned pasture, species richness, composition and relative abundance were monitored over a two year period both on and off artificially created earth mounds (100, 900, 8100 cm²). Mean species richness increased by up to two species per small mound (100 cm²) and by up to four species per large mound (8100 cm²). However, increased species richness was evident for less than two years. Initially, up to sixteen of the twenty species present occurred more often on earth mounds than off mounds, with two of these species found only on large mounds (8100 cm²). After two years, there was little or no significant difference in species composition and relative frequency on and off earth mounds. Experimental soil disturbance temporarily altered community structure simply by increasing space available for colonization since light, nutrient and water supply did not increase significantly on mounds. Soil disturbance can increase species richness and change species' relative frequency in disturbances as small as 100 cm² but such changes were likely too small and short lived to alter permanently the structure and rate of succession in the abandoned pasture studied here.     

Influence of vegetation and soil types on the wheatbelt termite,Drepanotermes tamminensis (Hill), in the Western Australian wheatbelt

A survey of the distribution and density of mounds of the harvester termite,Drepanotermes tamminensis (Hill), was carried out in the Durokoppin Nature Reserve, Western Australia in 1990. Vegetation and, to a lesser extent, soil type, appear to be important factors in determining density and distribution of termite mounds within the Reserve. A more detailed study of mounds in Wandoo (Eucalyptus capillosa) woodland and Casuarina (Allocasuarina campestris) shrubland indicated that the total number and size of mounds were significantly higher in the woodland than in the shrubland. The total wet weight biomass ofD. tamminensis was calculated as 3.74 gm⁻² (37.4 kg ha⁻¹) in the woodland and 1.69 gm⁻² (16.9 kg ha⁻¹) in the shrubland. Thus, of the two favored habitats, Wandoo woodland appears to be more optimal for this termite species than the Casuarina shrubland.     

Studies on a population of large termite mounds in Uganda

• 1 Two study-sites near Kampala were mapped and all of the termite mounds within them were measured. Changes were recorded in the three succeeding years. A method was developed for estimating mound volumes from height and diameter.
• 2 Macrotermes bellicosus mounds reached a volume of about 2.4 m³ in 3 years after which they grew more slowly to a maximum from 4 to 6 m³. Growth in volume during the first 3 years was approximately linear and its rate did not vary seasonally, probably because growth is related to temperature which was nearly constant throughout the year.
• 3 M.bellicosus mounds had an average length of life of about 10 years, although colonies survived on average only 4 years. The corresponding figures for Pseudacanthotermes spp. were 5 and 2 1/2 years. Some mounds were recolonized after the death of the original colony.
• 5 Fresh earth was brought up at a minimum rate of about 10 m³ ha^-1 yr^-1 but more was lost by erosion and destruction and the populations of mounds were decreasing.
• 4 The number of ventilation shafts in M.bellicosus mounds increased with the volume of the mound. Many mounds, both dead and alive, supported vegetation.
• 6 Local opinion is that Pseudacanthotermes are harmless but M.bellicosus is regarded as a pest and attempts are made to destroy its mounds.

     

Growth and demography ofCubitermes speciosus mounds (Isoptera,Termitidae)

The growth and demography ofCubitermes speciosus mounds were studied in an equatorial forest ecosystem near Kisangani, Zaïre. The mounds were censused and measured on eight occasions from June 1985 to January 1993 on 25 plots of 100 m².Mounds were found to grow in two ways: either they were enlarged sporadically, leading to a correlation between age and size; or they were no longer enlarged after a 2-year period of initial growth, which means that both large and small mounds can be old.The number of living mounds was fairly constant: it varied between 112 and 152 ha^–1 with an average of 139 ha^–1, and about a quarter of them were renewed each year. However, a clear tendency was observed over our 7.5-year observation period: the average size of the mounds increased regularly and was 2.7 times larger in 1993 than in 1985.Monthly birth and death rates were estimated using a model of continuous growth. Life expectancy at the age of 3 months was estimated from a survivorship curve at 3.5 years.     

Effects of ants on plant diversity in semi-natural grasslands

Ants (L. niger and L. flavus) build conspicuous mounds that are covered with vegetation. The aim of this study was to investigate whether the vegetation on ant mounds in semi-natural grasslands differed from that around the mounds. Another aim was to investigate whether the changes in the vegetation on ant mounds were influenced by grazing management or by habitat characteristics, semi-dry versus moist. Here, the total number of plant species and total plant cover were lower on ant mounds than in patches off-mound. The plant cover of perennials that form rosettes was twice as high on mounds inhabited by L. niger than on those inhabited by L. flavus. Only a few plant species were restricted to either ant mounds or adjacent field and the effects of ants on the plant diversity in semi-natural grasslands seemed to be low. Grazing management did not affect the differences in the vegetation on ant mounds and in equal-sized patches off-mound, whereas habitat characteristics affected ant-induced changes in vegetation cover of some plant species.     

Mima mounds in Kenya–a case of mistaken identity

Earth mounds at seven sites in upland areas of Kenya were examined. Termite activity was recorded from 91% of mounds and the major termite genus present was Odontotermes. Generally, signs of the mole rat, Tachyoryctes splendens were few and were present on only 18% mounds. All termite activity was mound-centred but only at Kiserian were mounds obvious centres for mole rat activity. At Molo, where Tachyoryctes were abundant, neither mounds nor much sign of termite activity were observed. The data are discussed in relation to the work of Cox and Gakahu. No convincing evidence was found to support the conclusion made by these workers for a mole rat origin of large earth mounds. The weight of the data suggests that a termite origin for these mounds is more likely.     

The influence of harvester and Messor capensis nest-mounds on the productivity and distribution of some plant species in the southern Karoo,South Africa

Nest-mounds of the harvester ant Messor capensis occur on and around nutrient-rich patches, along minor drainage lines in nutrient-rich soils, and on the plains, generally in nutrient-poor soils. Nest-site selection is related to the presence of suitable deep soils, the presence of stones and the distance from the nearest neighbouring nest. Two plant species, Galenia fruticosa and Pteronia pallens, were significantly associated with Messor capensis nest-mounds, both in numbers of mounds occupied and in numbers of individuals. A third species, Drosanthemum montaguense, was also more common in numbers of individuals, while a fourth species, Rhinephyllum macradenium was negatively associated with these mounds. The analysis of species guilds by soil type shows that significantly more species of nutrient-rich soils are present on M. capensis nest-mounds. Also, significantly more taller, woody species occurred on nest-mounds than in inter-mound spaces. Two species, Pteronia pallens and Osteospermum sinuatum, growing on ant nest-mounds had significantly longer inter-nodes than the same species growing off mounds. However, two other species, Pteronia cf. empetrifolia and Galenia fruticosa showed no difference in inter-node lengths between plants growing on and off mounds. Six of the nine species of plants sampled on ant nest-mounds had significantly higher seed production than plants of the same species growing in inter-mound spaces. The other three species showed a tendency towards more seeds per plant on ant nest-mounds. The proportions of live and dead plants on mounds differed between species. Only Ruschia spinosa showed a significant difference between the numbers of dead plants in the population on and off mounds, with more dead plants occurring on mounds. Significantly more seeds set on individuals of Pteronia pallens growing on ant nest-mounds than those growing off nest-mounds, but no such difference occurred in P. cf. empetrifolia. There was no significant difference in the proportion of seeds parasitized by the tephritid fly Desmella anceps for individuals of P. pallens and P. cf. empetrifolia growing on and off mounds.     

A close look at late carboniferous algal mounds: Schulterkofel,Carnic Alps,Austria

Summary During the uppermost Carboniferous and lowermost Permian algal mounds were formed in inner shelf settings of the Carnic Alps (Austria/Italy). A specific mound type, characterized by the dominance of the dasyclad green alga Anthracoporella was studied in detail with regard to geometry, relationship between mound and intermound rocks, composition of the sediment, biota and diagenetic criteria. The two meter-sized mounds studied, occur within depositional sequences of transgressive systems tracts in the Lower Pseudoschwagerina Limestones (uppermost Gzhelian) at the flank of the Schulterkofel. The mounds consist of an Anthracoporella core facies with a spongecrust boundstone facies at the base and at the top. The massive limestones of the Anthracoporella core facies exhibit abundant algal tufts and bushes, frequently in life position. The limestones of the intermound facies represented by thin-bedded bioclastic wackestones and packstones with abundant phylloid algae underlie and overlie the mounds. Intercalations of intermound beds within the mound facies indicate sporadic disruption of mound growth. Onlapping of intermound beds on steep mound flanks indicate rapid stabilization and lithification of mound flanks and the existence of a positive paleorelief. Asymmetrical shape of the mounds may be current controlled. Mound and intermound biota differ in the prevailing algae but are relatively similar with regard to associated foraminifera. Conspicuous differences concern bioerosion and biogenic encrustations. Bothare, high in intermound areas but low in the Anthracoporella core facies. The mounds show no ecological zonation. The mounds grew by in-place accumulation of disintegrated algal material and trapped bioclastic material between erect algal thalli. The comparison of the various Anthracoporella mounds demonstrates that almost each mound had ist own history. Establishing a general model for these mounds is a hazardous venture.     

First geo-marine survey of living cold-water Lophelia reefs in the Ionian Sea (Mediterranean basin)

Abstract Prosperous deep coral mounds including living colonies of Lophelia pertusa together with Madrepora oculata and Desmophyllum dianthus (= D. cristagalli) have been discovered in 2000, by fishery operations on the eastern side of the Ionian Sea. The living coral mounds are located between ca. 300 and 1,100 m on a gently dipping shelf off Apulia at Santa Maria di Leuca (SML), and characterized by a complex seabed topography. Side scan sonar, shallow high-resolution seismics and sampling indicate that these Lophelia-bearing coral mounds colonize quasi-indurate (firmground) Pleistocene sediment. At places live corals were found on Pleistocene coral-hardgrounds. The fauna associated with these Ionian modern coral mounds is less diversified than modern Eastern Atlantic counterparts. The core of living coral mounds colonies is at present located in 500–700 m and is tentatively suggested that their survival is mostly controlled by oceanographic factors. The SML coral banks represent so far a unique example of living Lophelia-bearing coral mounds in the Mediterranean basin.This revised version was published online in January 2005 with corrections to the author name and figure captions 4 and 5.     

Effect of three mound-building ant species on the formation of soil seed bank in mountain grassland

The effect of three ant species (Lasius flavus, Formica spp., Tetramorium caespitum) on soil seed bank formation was studied in temperate mountain grassland. Seed removal experiments, analysis of soil seed content and seed survival experiments were carried out to evaluate the influence of ground ants on the seed fate. In the seed removal experiment seeds of 16 species, including 5 species with elaiosome-bearing seeds (myrmecochores), were exposed and their removal followed for 39 h. On average, ants removed 63.8% of myrmecochorous seeds and 10.9% of seeds without adaptation to ant dispersal. Analysis of soil seed content revealed that myrmecochores, in spite of expectations that they would accumulate in nests of seed dispersing ants, were most abundant in the soil of control plots. Evidence on seed relocation to the ant nests was obtained from a comparison of mounds of seed dispersing and seed non-dispersing ant species, as more seeds were found in the mounds of Formica spp. and Tetramorium caespitum (seed dispersers) in comparison with the mounds of Lasius favus (non-disperser).The soil seed bank of the compared microhabitats (control plots and mounds of 3 ant species) differed in their species composition, seed abundance and vertical distribution. The most distinct qualitative differences were between seed flora of control plots and mounds of Tetramorium caespitum. Control plots had approximately 30,000 propagules per m², which was double the number of seeds found in the ant mounds. In control plots, abundance and diversity of seeds steeply declined with depth; this trend was not observed in the mounds probably due to bioturbation. In the seed survival experiment, more seeds (2 out of 3 species) survived in control plots, which may also contribute to the higher seed abundance in this microhabitat.This study showed that seed relocation by ants does not contribute significantly to seed bank build-up at this study site. Ants may, however, increase the regeneration success of myrmecochores, mainly by dispersal for distance and placement in a larger spectrum of microsites, in contrast to species not adapted for myrmecochory.     

A comparison of sites suitable for the seedling establishment of two co-occurring species, Swintonia glauca and Stemonurus scorpioides, in a tropical peat swamp forest

Spatial and temporal ground-surface dynamics are major factors that affect regeneration and species coexistence in tropical peat swamp forests. We studied the seedling survivorship and morphological features of two tree species that play important roles in maintaining the ground-surface dynamics of a peat swamp forest in Sumatra. Large Swintonia glauca trees form mounds, whereas large Stemonurus scorpioides trees occupy non-mounds. We monitored the demography of naturally dispersed Swintonia and Stemonurus seedlings that germinated in 2000. Survivorship of Swintonia seedlings was high under conditions of late germination, high-light environment, and elevated ground surface, and was negatively affected by distance to the nearest conspecific adult. Survivorship of Stemonurus was high under conditions of early germination and high conspecific seedling density, and was also negatively affected by distance to the nearest conspecific adult. The allometric features of Stemonurus seedlings indicated characteristics of stress tolerance, that is, low growth rate and thick, porous roots. Stemonurus, which has large wingless seeds, regenerated in non-mounds around the parental trees, while winged Swintonia seeds dispersed farther from the parent and established in patchily distributed gaps and mounds. Thus, Swintonia seedlings can survive on non-mound sites within gaps and possibly create mounds, while Stemonurus seedlings tend to maintain non-mounds around the parental trees.     

Effects of controlled livestock grazing and annual prescribed fire on epigeal termite mounds in a savannah woodland in Burkina Faso

The diversity and abundance of epigeal termite mounds were investigated in response to controlled livestock grazing and annual prescribed fire in a Sudanian savannah-woodland in Tiogo State Forest. Sampling of termite mounds was carried out in 4×4 subplots of 0.25 ha in a split-plot experimental design during the rainy season in 2002. There were two main plots of which one was fenced to exclude livestock grazing and the second exposed to grazing. Each of the main plots included 4 subplots with annual prescribed fire since 1992 and 4 subplots without fire. Data were collected on the number and characteristics of termite mounds. A mean density of 698 mounds ha⁻¹ was recorded. Mounds built by Trinervitermes spp. were the most abundant followed by Cubitermes spp., Macrotermes subhyalinus and M. bellicosus. The large mound-builders Macrotermes spp. dominated the community interms of basal area (96% of the total) and above-ground volume (99%). The diversity of mound types was notaffected by livestock grazing and annual early prescribed fire (P>0.05). There was no statistical effect of livestock grazing on mound density, whereas a strong depressive effect of annual fire was observed for Trinervitermes spp. mound density (P=0.012). In this ecosystem, annual prescribed fire appeared to be the major determinant for termite mound abundance. Received 2 February 2007; revised 23 October 2007 and 21 January 2008; accepted 22 Feburary 2008.     

Relative uptake of32P by cassava,banana, elephant foot yam and groundnut in intercropping systems

Absorption of applied³²P by the treated as well as neighbouring plants in two- and three-crop intercropping systems involving cassava (Manihot esculenta Crantz), banana (Musa (AAB) Mysore), elephant foot yam (Amorphophallus campanulatus Blume) and groundnut (Arachis hypogaea L.) was studied in field trials. Radiophosphorus applied to the root zone of one of the component species in the mixed systems was found to be absorbed not only by the treated plant but also by the neighbouring plants. Banana was the most dominant species in the cassava-banana-elephant foot yam intercropping system and accumulated the major portion of the radioactivity recovered in the whole system. Cassava planted on raised mounds absorbed³²P from the root zones of elephant foot yam and banana growing in the interspaces. Absorption of³²P from cassava mounds by elephant foot yam was negligible.In cassava-groundnut intercropping system, cassava was the dominant component accumulating about 96 to 99 per cent of the total³²P recovery in the system when the radiolabel was applied to cassava and about 48 to 88 per cent when applied to the intercrops depending on whether cassava was planted on paired row-ridge, mound or flat bed. The groundnut was able to absorb only negligible quantity of³²P from cassava root zone. The absorption of³²P by treated groundnut was highest in paired-row ridge method of planting and lowest in flat bed method of planting.     

Morphology and sedimentology of (clustered) cold-water coral mounds at the south Rockall Trough margins,NE Atlantic Ocean

Cold-water coral mounds on both margins of the Rockall Trough (NE Atlantic Ocean) have a strongly different morphology. Single, isolated mounds occur on the SE margin and are mainly found on the upper slope between 900 and 650 m water depth, while large mound clusters are found on the SW margin in water depths between 600 and 1,000 m, in a narrow zone almost parallel to the slope. Sedimentation rates on the mounds are higher than on the surrounding seabed as a result of baffling of biogenic carbonate debris and siliciclastic particles by the coral framework covering the mounds. This is confirmed by ²¹⁰Pb measurements. The individual coral growth rate can be three times higher then the vertical growth rate of the coral cover (±10 mm year⁻¹) which in turn is more than an order of magnitude higher then the present-day overall mound growth rate (±0.25 mm year⁻¹). The presence of extensive hardgrounds and firmgrounds and the three-dimensional coral framework are considered to be responsible for the stability of the relatively steep slopes of the mounds. High current velocities in the intramound areas result in local non-sedimentation and erosion, as is shown by the presence of IRD (ice-rafted debris) lag deposits on the seabed and moats around some of the mounds. The morphology and sedimentology of cold-water coral-covered (mainly Lophelia pertusa and Madrepora oculata) mounds on the southern Rockall Trough margins (NE Atlantic Ocean) is discussed and a model describing the development of these mounds is presented.     

Plant Colonizers Shape Early N-dynamics in Gopher-mounds

Disturbances by fossorial mammals are extremely common in many ecosystems, including the California annual grassland. We compared the impact of juveniles of four common plant colonizers (Aegilops triuncialis, Cerastium glomeratum, Aphanes occidentalis and Lupinus bicolor) on the pools and fluxes of N in mounds created by pocket gophers (Thomomys bottae Mewa). The mechanisms and magnitude of biotic N retention differed among plant species. In mounds colonized by Cerastium, Aphanes and Lupinus, the microbial N pool was significantly larger than the plant N pool, as is typical in California grasslands in the early spring, whereas in mounds colonized by Aegilops, there was a more equal distribution of biotic N between plant and microbial pools. A 1-day ¹⁵N pulse field experiment demonstrated that plant species significantly differed in their effects on the distribution of isotopic N, with the N-fixing Lupinus leaving most (82%) ¹⁵N as inorganic N in soil, whereas more ¹⁵N was immobilized in plants or otherwise removed from the available soil pool in mounds colonized by other species. The impacts of early colonizers on N dynamics suggest that the identity of plant species that initially colonize gopher mounds may have important consequences on the dynamics of the overall grassland community.     

Successional trends and biomass of mosses on windthrow mounds in the temperate rainforests of Southeast Alaska

We investigated successional trends on windthrow mounds in two old-growth Tsuga heterophylla-Picea sitchensis forests in northern southeast Alaska to determine the influence of windthrow disturbance on the maintenance of plant diversity. We were particularly interested in assessing the value of mosses in detecting long-term effects of disturbance in temperate rainforests. Mosses established a dense carpet on windthrow mounds within the first few decades after the disturbance. No consistent changes were noted in total moss and vascular plant cover, moss biomass, or species diversity between young mounds (±50 yrs), intermediate mounds (±150 yrs) or old mounds (> 200 yrs), or between mounds and the undisturbed forest floor, despite consistent differences in soils development.Classification and ordination of the vegetation data did not show a consistent relationship between soil surface age or soil depth and overall species composition on the two sites. Young mounds were the most compositionally distinctive, primarily due to moss species. Pogonatum alpinum var. sylvaticum, P. contortum and Polytrichum formosum were generally confined to young mounds with unstable substrata, while Dicranum majus and Sphagnum girgensohnii were associated with old soil surfaces and deep organic soils. Vascular plant species with affinities for riparian or deep shade habitats (Tiarella trifoliata, Coptis asplenifolia and Dryopteris expansa) showed a general preference for the forest floor. Gymnocarpium dryopteris was the only vascular plant with a significant association with young mounds.Mosses comprised approximately 25% of understory plant biomass and as much as 50% of understory productivity. In cool temperate forests, the inclusion of mosses in vegetation analysis may provide valuable insights into the nature of vegetation patterns over subtle environmental gradients. The distinctiveness of the temperate rainforest type and the unique ecological effects of windthrow disturbance in this type are also suggested by this study.Abbreviations HR = Heintzleman Ridge (study site) - OP = Outer Point (study site)     

Foraging strategy in a subterranean rodent,Spalax ehrenbergi: a test case for optimal foraging theory

Summary The foraging behavior of the subterranean mole rat Spalax ehrenbergi (Rodentia, Spalacidae) was tested according to the framework of optimal foraging theory. We compared the frequencies of food species hoarded in storage chambers of mole rats with the frequencies of these species occurring in the vicinity of the mole rats' nest mounds during the winter and spring seasons. In addition, we examined the food composition of several summer nest mounds. Laboratory observations were conducted in order to test the foraging behavior of mole rats under simulated subterranean conditions. The mole rat is a generalist and collects a variety of food species. Out of 33 plant species that were hoarded by mole rats in the 21 studied nest mounds, 61% (n=20) were geophytes, 21% (n=7) perennial herbs, 15% (n=5) annual herbs and 3% (n=1) dwarf shrubs. The frequency of each collected species in the 16 winter and spring nest mounds is in general accordance with its frequency in the mole rat's territory. This implies that the mole rat randomly samples the food reserve of its territory without special preference or directed search for a particular species. The collection or avoidance of any food item is not dependent on the presence or absence of any other food item. We suggest that the foragin generalism of the mole rat is a product of the constraints of a subterranean niche — the necessity to hoard food as much as possible in a limited time period and the high energetic investment of tunneling to the food items.     

Thicket clumps: A characteristic feature of the Kagera savanna landscape,East Africa

Question: What are the genesis and development of thicket clumps within a savanna landscape at geomorphically different locations and what are the driving forces? Location: The Kagera Region, in the border area of Rwanda, Uganda and Tanzania. Methods: The vegetation of 32 dry evergreen thicket clumps and their surrounding savannas have been analysed at different geomorphic locations. At each vegetation plot Na⁺, K⁺, Mg²⁺, Ca²⁺, Al³⁺, Fe^2+/3+, H⁺, P, C, N, bulk density and particle size were determined for each soil horizon. The impact of soil and termite mounds on thicket clump dynamics on seasonally waterlogged plains, gentle slopes and stony hillsides were assessed. Results: Thicket clumps and their surrounding savannas have a distinct structure and floristic composition. They also have distinct soil properties although parent materials are the same. On seasonally waterlogged plains, new thicket clumps can develop on Macrotermitinae mounds; on stony hillsides, Trinervitermes and Macrotermes show a uniform distribution pattern and may initiate the genesis of thicket clumps. Conclusions: Geomorphology broadly determines the significance and interactions of the main factors affecting site‐specific vegetation dynamics. On seasonally waterlogged plains, thicket clumps are restricted to termite mounds. Since intra‐species competition dictates a minimal distance between neighbouring Macrotermitinae colonies, thicket clumps do not coalescence. By contrast, on stony hillsides, the vegetation mosaic is highly dynamic and determined by the interplay of several factors. The growth of thicket clumps is mainly a function of the fire regime and the browsing intensity. At the present time, frequent cool, early dry season fires and the near absence of large browsers have favoured the advance and coalescence of thicket clumps and forest patches on stony hillsides.     

The dynamics of a succulent karoo vegetation

Strong inference techniques were used in a preliminary study of the structure and dynamics of the vegetation the Prince Albert region of the Great Karoo, South Africa. Seedling emergence studies indicate that open areas in the interstices of the existent vegetation are colonized by mound-building species of the Mesembryanthemaceae. Later, these species serve as sites of establishment for seedlings of several species of woody shrubs. Eventually, the woody shrubs replace the mound-building mesems through interspecific competition. The woody shrubs persist in the community until they reach senescence and die or are removed through overgrazing. Superimposed on this dynamic pattern is a further temporal pattern involving a combination of disturbance and subsequent soil changes. Much of this effect is caused by fossorial animals, which are associated with large circular (10–20 m diameter) mounds. Burrowing by these animals changes the soil characteristics and establishes an additional sequence of vegetation succession in which the mounds serve as refuges from which non-palatable species, such as Pteronia pallens, can invade the rangeland. Later, as the mounds are abandoned and disturbance effects cease, more palatable species, such as P. empetrifolia, will replace P. pallens, overcoming finally the effects of the adjacent mounds on the surrounding vegetation.